Lophiosilurus é um gênero monotípico representado por L. alexandri, uma espécie endêmica da bacia do rio São Francisco, Brasil. Em análises filogenéticas anteriores, o gênero foi recuperado como grupo irmão de Cephalosilurus. No entanto, poucas espécies de Cephalosilurus ou poucos caracteres foram incluídos nesses estudos. Assim, este estudo tem como objetivo testar a hipótese de monofilia dos gêneros Lophiosilurus e Cephalosilurus, com uma análise filogenética mais abrangente, incluindo todas as espécies de Cephalosilurus e um número representativo de caracteres. As análises filogenéticas de 18 táxons terminais (15 do grupo interno e três grupos externos) foram realizadas com base em uma matriz combinada de 75 caracteres, incluindo 70 caracteres morfológicos discretos de osteologia e neuroanatomia, quatro caracteres contínuos e um de morfometria geométrica da cabeça. A monofilia da família Pseudopimelodidae foi altamente apoiada e Cephalosilurus foi sinonimizado como Lophiosilurus. A filogenia recuperada do gênero foi (L. albomarginatus (L. nigricaudus (L. apurensis (L. fowleri, L. alexandri))).

Palavras-chave:
Biodiversidade; Cephalosilurus; Pacamã; Sistemática; Taxonomia

Autoria

Oscar A. Shibatta

Programa de Pós-Graduação em Ciências Biológicas, Departamento de Biologia Animal e Vegetal, Centro de Ciências Biológicas, Universidade Estadual de Londrina, 86057-970 Londrina, PR, Brazil. (OAS) shibatta@uel.br (corresponding author); (LSS) lenicesouza@hotmail.com.Universidade Estadual de LondrinaBrazilLondrina, PR, BrazilPrograma de Pós-Graduação em Ciências Biológicas, Departamento de Biologia Animal e Vegetal, Centro de Ciências Biológicas, Universidade Estadual de Londrina, 86057-970 Londrina, PR, Brazil. (OAS) shibatta@uel.br (corresponding author); (LSS) lenicesouza@hotmail.com.

http://orcid.org/0000-0003-2883-5934

Lucas R. Jarduli

Centro Universitário das Faculdades Integradas de Ourinhos, 19909-100 Ourinhos, SP, Brazil. (LRJ) lucasjarduli@gmail.com.Centro Universitário das Faculdades Integradas de OurinhosBrazilOurinhos, SP, BrazilCentro Universitário das Faculdades Integradas de Ourinhos, 19909-100 Ourinhos, SP, Brazil. (LRJ) lucasjarduli@gmail.com.

http://orcid.org/0000-0002-5796-6434

Vitor P. Abrahão

Programa de Pós-Graduação em Biodiversidade e Evolução, Instituto de Biologia, Universidade Federal da Bahia, Rua Barão de Geremoabo, 668, Ondina, 40170-115 Salvador, BA, Brazil. (VPA) vitorabrahao32@gmail.com.Universidade Federal da BahiaBrazilSalvador, BA, BrazilPrograma de Pós-Graduação em Biodiversidade e Evolução, Instituto de Biologia, Universidade Federal da Bahia, Rua Barão de Geremoabo, 668, Ondina, 40170-115 Salvador, BA, Brazil. (VPA) vitorabrahao32@gmail.com.

http://orcid.org/0000-0002-6596-5079

Lenice Souza-Shibatta

http://orcid.org/0000-0001-8097-6895

Edited-by

Gloria Arratia

Correspondence

Oscar A. Shibatta shibatta@uel.br

Author's Contribution

Oscar Akio Shibatta: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Supervision, Validation, Visualization, Writing-original draft, Writing-review and editing.

Lucas Ribeiro Jarduli: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Validation, Visualization, Writing-original draft, Writing-review and editing.

Vitor Pimenta Abrahão: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Validation, Visualization, Writing-original draft, Writing-review and editing.

Lenice Souza-Shibatta: Conceptualization, Formal analysis, Investigation, Methodology, Validation, Visualization, Writing-original draft, Writing-review and editing.

Competing Interests

The authors declare no competing interest

Ethical Statement

Specimens examined in this study were available in collections.

SCIMAGO INSTITUTIONS RANKINGS

Figuras | Tabelas

Figuras (12)
Tabelas (1)

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FIGURE 1 |
Geographic distribution of Lophiosilurus species. Lophiosilurus apurensis, MZUEL 6492, 189.7 mm SL, Apure river basin, Venezuela; L. nigricaudus, INPA 21632, 58.1 mm SL, Sipaliwini river basin, Suriname; L. albomarginatus, ROM 61336, 88.4 mm SL, Tukeit river basin, Guyana; L. alexandri, MZUEL 5377, 157.8 mm SL, São Francisco river basin, Brazil; L. fowleri, FMNH 54254, holotype, 301.4 mm SL, São Francisco river basin, Brazil.

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FIGURE 2 |
Previous hypothesis of Pseudopimelodidae phylogeny: A.Shibatta, (2003b)Shibatta OA. Family Pseudopimelodidae (Bumblebee catfishes, dwarf marbled catfishes). In: Reis RE, Kullander SO, Ferraris Jr CJ, organizers. Check List of the Freshwater Fishes of South and Central America. Porto Alegre: Edipucrs; 2003b. p.401–405.; B.Ortega-Lara, Lehmann, (2006)Ortega-Lara A, Lehmann PA. Cruciglanis, a new genus of Pseudopimelodid catfish (Ostariophysi: Siluriformes) with description of a new species from the Colombian Pacific coast. Neotrop Ichthyol. 2006; 4(2):147–56. https://doi.org/10.1590/S1679-62252006000200002
https://doi.org/10.1590/S1679-6225200600... ; C.Birindelli, Shibatta, (2011)Birindelli JLO, Shibatta OA. Morphology of the gas bladder in bumblebee catfishes (Siluriformes, Pseudopimelodidae). J Morphol. 2011; 272(7):890–96. https://doi.org/10.1002/jmor.10961
https://doi.org/10.1002/jmor.10961... ; D.Sullivan et al., (2013)Sullivan JP, Muriel-Cunha J, Lundberg JG. Phylogenetic relationships and molecular dating of the major groups of catfishes of the Neotropical superfamily Pimelodoidea (Teleostei, Siluriformes). P Acad Nat Sci Phila. 2013; 162:89–110. https://www.jstor.org/stable/42751950
https://www.jstor.org/stable/42751950... ; E.Shibatta, Vari, (2017)Shibatta OA, Vari RP. A new genus of Neotropical rheophilic catfishes, with four new species (Teleostei: Siluriformes: Pseudopimelodidae). Neotrop Ichthyol. 2017; 15(2):e160132. https://doi.org/10.1590/1982-0224-20160132
https://doi.org/10.1590/1982-0224-201601... ; F.Rangel-Medrano et al., (2020)Rangel-Medrano J, Ortega-Lara A, Márquez EJ. Ancient genetic divergence in bumblebee catfish genus Pseudopimelodus (Pseudopimelodidae: Siluriformes) from northwestern South America. PeerJ. 2020; 8:e9028. http://doi.org/10.7717/peerj.9028
http://doi.org/10.7717/peerj.9028... ; G.Silva et al., (2021)Silva GSC, Melo B, Roxo FF, Ochoa LE, Shibatta OA, Sabaj MH, Oliveira C. Phylogenomics of the bumblebee catfishes (Siluriformes: Pseudopimelodidae) using ultraconserved elements. Jour Zool Syst Evol Res. Forthcoming 2021. http://doi.org/10.1111/jzs.12513
http://doi.org/10.1111/jzs.12513... .

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FIGURE 3 |
Landmark position on Pseudopimelodus mangurus, MZUEL 1073, 185.2 mm SL. 0 = Tip of snout; 1 = Posterior margin of the left anterior nostril; 2= Posterior margin of the right anterior nostril; 3 = Insertion of left maxillary barbel; 4 = Insertion of right maxillary barbel; 5 = Posterior margin of the left posterior nostril; 6 = Posterior margin of the right posterior nostril; 7 = Medial margin of left eye; 8 = Medial margin of right eye; 9 = Most lateral point of the head in the left; 10 = Most lateral point of the head in the right; 11 = Upper end of the left operculum; 12 = Upper end of the right operculum; 13 = Insertion of dorsal fin spine.

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FIGURE 4 |
Dorsal view of left maxillary bone; anterior up: A. Steindachneridion scriptum, MZUEL 19302; B. Rhamdia quelen, MZUEL 14631; C. Lophiosilurus apurensis, MBUCV-V-15379. Scale bars = 1 mm.

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FIGURE 5 |
Neurocranium in dorsal view: A. Lophiosilurus alexandri, MZUEL 5542; B. L. fowleri, MZUEL 12095; C. L. apurensis, MBUCV-V-15379; D. L. albomarginatus, ROM 61336; E. Batrochoglanis raninus, MZUSP 23407; F. Microglanis parahybae, MNRJ 15989; G. Pseudopimelodus mangurus, MZUEL 12937; H. Rhyacoglanis paranensis, MZUEL 6034. Bones in yellow and cartilage in bluish grey. Scale bars = 5 mm.

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FIGURE 6 |
Ventral view of left ectopterygoid and entopterygoid bones; anterior up: A. Batrochoglanis raninus, MZUSP 23407; B. Lophiosilurus albomarginatus, ROM 61336; C. Lophiosilurus alexandri, MZUEL 5542.

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FIGURE 7 |
Shape of gill rakers on the first ceratobranchial; lateral view of left selected portion; anterior to left: A. Steindachneridion scriptum, MZUEL 19302; B. Lophiosilurus alexandri, MZUEL 5542. Scale bars = 5 mm.

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FIGURE 8 |
Ventral view of complex vertebrae; anterior up: A. Ictalurus punctatus, MZUEL 13834; B. Steindachneridion scriptum, MZUEL, 19302; C. Pseudopimelodus mangurus, MZUEL 17286; D. Lophiosilurus alexandri, MZUEL 5542. Scale bar = 5 mm.

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FIGURE 9 |
Lateral view of caudal bones; anterior to left: A. Steindachneridion scriptum, MZUEL 19302; B. Rhamdia quelen, MZUEL 11664; C. Pseudopimelodus mangurus, MZUEL 17286; D. Lophiosilurus alexandri, MZUEL 13288. Scale bars = 5 mm.

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FIGURE 10 |
Dorsal view of brain; anterior up: A. Rhamdia quelen, MZUEL 6036, 222.4 mm SL; B. Rhyacoglanis paranensis, MZUEL 6034, 39.2 mm SL; C. Cruciglanis pacifici, MCP non-catalogued, 142.4 mm SL; D. Lophiosilurus alexandri, MZUEL 5377, 58.7 mm SL.

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FIGURE 11 |
Inferred phylogeny and landmark mapping (numbers) on the head of Lophiosilurus species. The arrow indicates the ancestral configuration. Enlarged image of Lophiosilurus alexandri in the box (grey lines = ancestral shape; red lines = reconstructed shape, blue lines = landmark position changing to the corresponding node in the ancestor).

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FIGURE 12 |
Resampled tree (CI = 63; RI = 76; length (best score) = 148.87) of Pseudopimelodidae derived from combined 75 characters matrix data set, with synapomorphies above and support values (symmetric resampling) under the branches. Pseudopimelodidae subfamilies names according to Silva et al., (2021)Silva GSC, Melo B, Roxo FF, Ochoa LE, Shibatta OA, Sabaj MH, Oliveira C. Phylogenomics of the bumblebee catfishes (Siluriformes: Pseudopimelodidae) using ultraconserved elements. Jour Zool Syst Evol Res. Forthcoming 2021. http://doi.org/10.1111/jzs.12513
http://doi.org/10.1111/jzs.12513... .

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TABLE 1 |
Key taxonomic publications with changes in the names of Lophiosilurus species. † undescribed fossil species identified by Lundberg et al., (2010)Lundberg JG, Sabaj MH, Dahdul WM, Aguilera OA. The Amazonian Neogene fish fauna. In: Hoorn C, Wesselingh FP, editors. Amazônia: Landscape and species evolution: A look into de past. Hoboken, NY: Blackwell Publishing Ltd.; 2010. p.281–301..

FIGURE 1 | Geographic distribution of Lophiosilurus species. Lophiosilurus apurensis, MZUEL 6492, 189.7 mm SL, Apure river basin, Venezuela; L. nigricaudus, INPA 21632, 58.1 mm SL, Sipaliwini river basin, Suriname; L. albomarginatus, ROM 61336, 88.4 mm SL, Tukeit river basin, Guyana; L. alexandri, MZUEL 5377, 157.8 mm SL, São Francisco river basin, Brazil; L. fowleri, FMNH 54254, holotype, 301.4 mm SL, São Francisco river basin, Brazil.

FIGURE 2 | Previous hypothesis of Pseudopimelodidae phylogeny: A.Shibatta, (2003b)Shibatta OA. Family Pseudopimelodidae (Bumblebee catfishes, dwarf marbled catfishes). In: Reis RE, Kullander SO, Ferraris Jr CJ, organizers. Check List of the Freshwater Fishes of South and Central America. Porto Alegre: Edipucrs; 2003b. p.401–405.; B.Ortega-Lara, Lehmann, (2006)Ortega-Lara A, Lehmann PA. Cruciglanis, a new genus of Pseudopimelodid catfish (Ostariophysi: Siluriformes) with description of a new species from the Colombian Pacific coast. Neotrop Ichthyol. 2006; 4(2):147–56. https://doi.org/10.1590/S1679-62252006000200002
https://doi.org/10.1590/S1679-6225200600... ; C.Birindelli, Shibatta, (2011)Birindelli JLO, Shibatta OA. Morphology of the gas bladder in bumblebee catfishes (Siluriformes, Pseudopimelodidae). J Morphol. 2011; 272(7):890–96. https://doi.org/10.1002/jmor.10961
https://doi.org/10.1002/jmor.10961... ; D.Sullivan et al., (2013)Sullivan JP, Muriel-Cunha J, Lundberg JG. Phylogenetic relationships and molecular dating of the major groups of catfishes of the Neotropical superfamily Pimelodoidea (Teleostei, Siluriformes). P Acad Nat Sci Phila. 2013; 162:89–110. https://www.jstor.org/stable/42751950
https://www.jstor.org/stable/42751950... ; E.Shibatta, Vari, (2017)Shibatta OA, Vari RP. A new genus of Neotropical rheophilic catfishes, with four new species (Teleostei: Siluriformes: Pseudopimelodidae). Neotrop Ichthyol. 2017; 15(2):e160132. https://doi.org/10.1590/1982-0224-20160132
https://doi.org/10.1590/1982-0224-201601... ; F.Rangel-Medrano et al., (2020)Rangel-Medrano J, Ortega-Lara A, Márquez EJ. Ancient genetic divergence in bumblebee catfish genus Pseudopimelodus (Pseudopimelodidae: Siluriformes) from northwestern South America. PeerJ. 2020; 8:e9028. http://doi.org/10.7717/peerj.9028
http://doi.org/10.7717/peerj.9028... ; G.Silva et al., (2021)Silva GSC, Melo B, Roxo FF, Ochoa LE, Shibatta OA, Sabaj MH, Oliveira C. Phylogenomics of the bumblebee catfishes (Siluriformes: Pseudopimelodidae) using ultraconserved elements. Jour Zool Syst Evol Res. Forthcoming 2021. http://doi.org/10.1111/jzs.12513
http://doi.org/10.1111/jzs.12513... .

FIGURE 3 | Landmark position on Pseudopimelodus mangurus, MZUEL 1073, 185.2 mm SL. 0 = Tip of snout; 1 = Posterior margin of the left anterior nostril; 2= Posterior margin of the right anterior nostril; 3 = Insertion of left maxillary barbel; 4 = Insertion of right maxillary barbel; 5 = Posterior margin of the left posterior nostril; 6 = Posterior margin of the right posterior nostril; 7 = Medial margin of left eye; 8 = Medial margin of right eye; 9 = Most lateral point of the head in the left; 10 = Most lateral point of the head in the right; 11 = Upper end of the left operculum; 12 = Upper end of the right operculum; 13 = Insertion of dorsal fin spine.

FIGURE 4 | Dorsal view of left maxillary bone; anterior up: A. Steindachneridion scriptum, MZUEL 19302; B. Rhamdia quelen, MZUEL 14631; C. Lophiosilurus apurensis, MBUCV-V-15379. Scale bars = 1 mm.

FIGURE 5 | Neurocranium in dorsal view: A. Lophiosilurus alexandri, MZUEL 5542; B. L. fowleri, MZUEL 12095; C. L. apurensis, MBUCV-V-15379; D. L. albomarginatus, ROM 61336; E. Batrochoglanis raninus, MZUSP 23407; F. Microglanis parahybae, MNRJ 15989; G. Pseudopimelodus mangurus, MZUEL 12937; H. Rhyacoglanis paranensis, MZUEL 6034. Bones in yellow and cartilage in bluish grey. Scale bars = 5 mm.

FIGURE 6 | Ventral view of left ectopterygoid and entopterygoid bones; anterior up: A. Batrochoglanis raninus, MZUSP 23407; B. Lophiosilurus albomarginatus, ROM 61336; C. Lophiosilurus alexandri, MZUEL 5542.

FIGURE 7 | Shape of gill rakers on the first ceratobranchial; lateral view of left selected portion; anterior to left: A. Steindachneridion scriptum, MZUEL 19302; B. Lophiosilurus alexandri, MZUEL 5542. Scale bars = 5 mm.

FIGURE 8 | Ventral view of complex vertebrae; anterior up: A. Ictalurus punctatus, MZUEL 13834; B. Steindachneridion scriptum, MZUEL, 19302; C. Pseudopimelodus mangurus, MZUEL 17286; D. Lophiosilurus alexandri, MZUEL 5542. Scale bar = 5 mm.

FIGURE 9 | Lateral view of caudal bones; anterior to left: A. Steindachneridion scriptum, MZUEL 19302; B. Rhamdia quelen, MZUEL 11664; C. Pseudopimelodus mangurus, MZUEL 17286; D. Lophiosilurus alexandri, MZUEL 13288. Scale bars = 5 mm.

FIGURE 10 | Dorsal view of brain; anterior up: A. Rhamdia quelen, MZUEL 6036, 222.4 mm SL; B. Rhyacoglanis paranensis, MZUEL 6034, 39.2 mm SL; C. Cruciglanis pacifici, MCP non-catalogued, 142.4 mm SL; D. Lophiosilurus alexandri, MZUEL 5377, 58.7 mm SL.

FIGURE 11 | Inferred phylogeny and landmark mapping (numbers) on the head of Lophiosilurus species. The arrow indicates the ancestral configuration. Enlarged image of Lophiosilurus alexandri in the box (grey lines = ancestral shape; red lines = reconstructed shape, blue lines = landmark position changing to the corresponding node in the ancestor).

FIGURE 12 | Resampled tree (CI = 63; RI = 76; length (best score) = 148.87) of Pseudopimelodidae derived from combined 75 characters matrix data set, with synapomorphies above and support values (symmetric resampling) under the branches. Pseudopimelodidae subfamilies names according to **Silva et al., (2021)**Silva GSC, Melo B, Roxo FF, Ochoa LE, Shibatta OA, Sabaj MH, Oliveira C. Phylogenomics of the bumblebee catfishes (Siluriformes: Pseudopimelodidae) using ultraconserved elements. Jour Zool Syst Evol Res. Forthcoming 2021. http://doi.org/10.1111/jzs.12513
http://doi.org/10.1111/jzs.12513... .

TABLE 1 | Key taxonomic publications with changes in the names of Lophiosilurus species. † undescribed fossil species identified by **Lundberg et al., (2010)**Lundberg JG, Sabaj MH, Dahdul WM, Aguilera OA. The Amazonian Neogene fish fauna. In: Hoorn C, Wesselingh FP, editors. Amazônia: Landscape and species evolution: A look into de past. Hoboken, NY: Blackwell Publishing Ltd.; 2010. p.281–301..

Nominal species	Mees (1974)	Shibatta (2003b)	Current contribution
Cephalosilurus fowleri Haseman, 1911	Pseudopimelodus fowleri	Cephalosilurus fowleri	Lophiosilurus fowleri (Haseman, 1911)
Cf. Cephalosilurus sp.†	‒	‒	Lophiosilurus sp.
Lophiosilurus alexandri Steindachner, 1876	‒	Lophsiosilurus alexandri	Lophiosilurus alexandri Steindachner, 1876
Pseudopimelodus albomarginatus Eigenmann, 1912	Pseudopimelodus albomarginatus	Cephalosilurus albomarginatus	Lophiosilurus albomarginatus (Eigenmann, 1912)
Pseudopimelodus apurensis Mees, 1978	‒	Cephalosilurus apurensis	Lophiosilurus apurensis (Mees, 1978)
Pseudopimelodus nigricauda Mees, 1974	Pseudopimelodus nigricauda	Cephalosilurus nigricaudus	Lophiosilurus nigricaudus (Mees, 1974)

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[1] Edited-by

Gloria Arratia